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Comparing Regional and Supra-Regional Transfer Functions for Palaeohydrological Reconstruction from Holocene Peatlands

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Comparing Regional and Supra-Regional Transfer Functions for Palaeohydrological Reconstruction from Holocene Peatlands promoting access to White Rose research papers Universities of Leeds, Sheffield and York http://eprints.whiterose.ac.uk/ This is an author produced version of a paper published in Palaeogeography, Palaeoclimatology, Palaeoecology. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/75724/ Published paper: Turner, TE, Swindles, GT, Blundell, A and Charman, DJ (2013) Comparing regional and supra-regional transfer functions for palaeohydrological reconstruction from Holocene peatlands. Palaeogeography, Palaeoclimatology, Palaeoecology, 369. 395 - 408. http://dx.doi.org/10.1016/j.palaeo.2012.11.005 White Rose Research Online [email protected] Comparing regional and supra-regional transfer functions for palaeohydrological reconstruction from Holocene peatlands 1T. Edward Turner, 1Graeme T. Swindles, 2Dan J. Charman, 1Antony Blundell 1School of Geography, University of Leeds, Leeds LS2 9JT, UK 2Department of Geography, University of Exeter, Exeter EX4 4RJ Corresponding author: E-mail: [email protected], Tel: +44 (0)113 34 31593 1 Abstract Testate amoebae-based transfer functions are commonly used in peatland palaeoclimate studies. These models have been developed in several regions of the world and are sometimes used for palaeohydrological reconstruction from fossil data in locations where no transfer functions exist. Limitations of this approach may include missing modern analogues and problems associated with site-specific or regional factors in testate amoebae ecology and biogeography. This study presents new testate amoebae-hydrology transfer functions based on data from six peatlands in Northern England. Transfer functions were generated for water table depth and moisture content using weighted averaging tolerance downweighted regression with inverse deshrinking and model performance was assessed using leave-one-out (jacknifing) cross-validation. To examine the robustness of applying transfer functions extra-regionally, we performed a number of spatially independent cross-validation (SICV) tests using contemporary testate amoebae and environmental data from established Northern Ireland and European transfer functions. Inferred water table depths were consistent with observed values in terms of position along the hydrological gradient, however magnitudes varied. We then applied the three independent transfer functions to fossil data from a peatland in Northern England to compare the reconstructions. The results show that the direction of the reconstructions is consistent in terms of wet/dry shifts. However, variation in the magnitudes of the reconstructed water tables is apparent. This probably reflects the sampling regime, including temporal/seasonal effects, and differences in testate amoebae ecology between regions and individual sites. Keywords: Peatlands, testate amoebae, transfer function, palaeoclimate, Holocene, cross-validation 2 1. Introduction The utilisation of transfer functions for quantitative palaeohydrological reconstruction from peatlands is now commonplace in several regions of the world (e.g. Booth, 2008; Charman, 1997; Lamentowicz et al., 2009; Lamentowicz et al., 2010; Langdon and Barber, 2005; Mitchell et al., 2001; Payne et al., 2006; Swindles et al., 2010). The hydrology of ombrotrophic (‘rain-fed’) peatlands is strongly linked to effective precipitation (precipitation – evapotranspiration), especially during the deficit period (Charman, 2007). Thus, inferences from peatland proxy-based palaeohydrological reconstructions contribute significantly to understanding Holocene hydroclimatic change. Although there are a number of peat-based hydrological proxies, such as plant macrofossils and peat humification, testate amoebae have become increasingly favoured by many researchers over the last two decades as transfer functions can be used for quantitative reconstruction of palaeo-water tables (e.g. Charman et al., 2007; Charman and Warner, 1992; Lamentowicz et al., 2010; Payne and Mitchell, 2007; Swindles et al., 2010; Wilmshurst et al., 2003; Woodland et al., 1998). Testate amoebae are a polyphyletic group of single-celled organisms that form a shell, or test. They are abundant on bog surfaces, especially living within the water film of bryophytes, and on death their empty tests are preserved in accumulating peat (Charman, 2001; Smith et al., 2008; Swindles and Roe, 2007; Warner, 1990). Testate amoebae form a significant part of the microbial ecosystem in peatland soils, enhancing nutrient cycles by consuming bacteria, protozoa, micro-algae, fungi and micrometazoa (Coûteaux and Darbyshire, 1998; Foissner, 1999; Lamentowicz and Mitchell, 2005; Mitchell et al., 2008a; Mitchell et al., 2000a). Community composition of testate amoebae is influenced by a number of biotic and abiotic factors and they respond 3 rapidly to environmental change (Buttler et al., 1996; Mitchell et al., 2000a; Tolonen, 1986). The dominant control on testate amoebae in peatlands has repeatedly been shown to be bog surface wetness (water table depth, moisture content) particularly in ombrotrophic systems where pH and nutrient status has a limited range (Bobrov et al., 1999; Booth, 2001; Charman, 2001; Lamentowicz and Mitchell, 2005; Payne et al., 2006; Swindles et al., 2009; Woodland et al., 1998). Although the position of testate amoebae in the microbial network is complex, their sensitivity to hydrology makes them ideal biological indicator organisms in both contemporary and palaeoecological studies (Turner and Swindles, 2012). The body of work on the ecology and biogeography of testate amoebae is extensive. Testate amoebae are often considered to be largely (with a number of exceptions) cosmopolitan in their distribution in Sphagnum-dominated peatlands, particularly in the Northern Hemisphere (Mitchell et al., 2000a; Smith et al., 2008). Testate amoebae- based hydrological transfer functions have been developed in many parts of the world; Alaska (Markel et al., 2010; Payne et al., 2006), Canada (Charman and Warner, 1997; Warner and Charman, 1994), Europe (Charman et al., 2007), Greece (Payne and Mitchell, 2007), New Zealand (Charman, 1997; Wilmshurst et al., 2003), North America (Booth, 2008; Booth and Zygmunt, 2005), Northern Ireland (Swindles et al., 2009), Switzerland (Mitchell et al., 1999), Poland (Lamentowicz et al., 2008; Lamentowicz and Mitchell, 2005), and the United Kingdom (Woodland et al., 1998). Due to spatial autocorrelation (Borcard et al., 1992; Lennon, 2000), transfer functions require validation against data sets that are completely independent of the training set (Belyea, 2007; Birks et al., 2010). However, very few models are rigorously tested using 4 spatially independent data and it has been suggested that they may generate reconstructions of unrealistic accuracy (Payne et al., 2011; Telford and Birks, 2005). Whilst the environmental preferences of these organisms are generally consistent between regions, their sensitivity to microsite variations and localised idiosyncrasies (Mitchell et al., 2000b) suggest it would be unreasonable to make palaeoecological inferences for one region using contemporary data from another (Charman et al., 2000). Therefore, a comprehensive understanding of the local ecology of testate amoebae is ideally a prerequisite when taking a transfer function approach to palaeoenvironmental reconstruction in any given area. To date, information on testate amoebae species- environment relationships in Britain comprise the dataset of Woodland et al. (1998) (163 samples in 9 sites) and 18 samples from a single British site included in the European transfer function of Charman et al. (2007). A need for further contemporary data from Britain is pertinent when considering the plethora of peatland palaeohydrological reconstructions from the region (e.g. Barber and Langdon, 2007; Blundell and Barber, 2005; Charman et al., 2004; Charman et al., 1999; Daley and Barber, in press; Hendon and Charman, 2004; Langdon and Barber, 2005). However, producing single-site training sets for each individual investigation would be impractical, and therefore the complexity of applying training sets extra-regionally needs further consideration. The aims of this study are to 1) examine environmental controls on contemporary testate amoebae communities in the Northern England region; 2) develop local transfer functions for quantitative palaeohydrological reconstruction based on fossil testate amoebae assemblages from the area; 3) to explore the issue of spatial scale and biogeography in transfer function development by comparing the results to recently 5 published regional (Swindles et al., 2009) and European (Charman et al., 2007) transfer functions; 4) to take a novel approach to model validation, using spatially independent data to rigorously test the potential of employing transfer functions extra-regionally. 2. Methods 2.1 Study Sites Six sites located in North and West Yorkshire, United Kingdom (Fig.1) were selected for this study to encompass a range of peatland environments within a similar climatic regime. Although the majority of palaeohydrological studies have been carried out on ombrotrophic raised bogs, capturing taxa associated with other bog types may be critical to understanding bog development trajectories. We included a variety of bog types in site selection to 1) ensure the inclusion of as many modern analogues as possible; and 2) sample full environmental gradients as recommended by Payne et al. (2011). However, we acknowledge that this sampling strategy cannot
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